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Charles Darwin University The extinction risk of New Zealand chondrichthyans Finucci, Brittany; Duffy, Clinton A.J.; Francis, Malcolm P.; Gibson, Claudine; Kyne, Peter M. Published in: Aquatic Conservation: Marine and Freshwater Ecosystems DOI: 10.1002/aqc.3053 Published: 01/05/2019 Document Version Peer reviewed version Link to publication Citation for published version (APA): Finucci, B., Duffy, C. A. J., Francis, M. P., Gibson, C., & Kyne, P. M. (2019). The extinction risk of New Zealand chondrichthyans. Aquatic Conservation: Marine and Freshwater Ecosystems, 29(5), 783-797. https://doi.org/10.1002/aqc.3053 General rights Copyright and moral rights for the publications made accessible in the public portal are retained by the authors and/or other copyright owners and it is a condition of accessing publications that users recognise and abide by the legal requirements associated with these rights. • Users may download and print one copy of any publication from the public portal for the purpose of private study or research. • You may not further distribute the material or use it for any profit-making activity or commercial gain • You may freely distribute the URL identifying the publication in the public portal Take down policy If you believe that this document breaches copyright please contact us providing details, and we will remove access to the work immediately and investigate your claim. Download date: 10. Apr. 2022

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Page 1: Charles Darwin University The extinction risk of New

Charles Darwin University

The extinction risk of New Zealand chondrichthyans

Finucci, Brittany; Duffy, Clinton A.J.; Francis, Malcolm P.; Gibson, Claudine; Kyne, Peter M.

Published in:Aquatic Conservation: Marine and Freshwater Ecosystems

DOI:10.1002/aqc.3053

Published: 01/05/2019

Document VersionPeer reviewed version

Link to publication

Citation for published version (APA):Finucci, B., Duffy, C. A. J., Francis, M. P., Gibson, C., & Kyne, P. M. (2019). The extinction risk of New Zealandchondrichthyans. Aquatic Conservation: Marine and Freshwater Ecosystems, 29(5), 783-797.https://doi.org/10.1002/aqc.3053

General rightsCopyright and moral rights for the publications made accessible in the public portal are retained by the authors and/or other copyright ownersand it is a condition of accessing publications that users recognise and abide by the legal requirements associated with these rights.

• Users may download and print one copy of any publication from the public portal for the purpose of private study or research. • You may not further distribute the material or use it for any profit-making activity or commercial gain • You may freely distribute the URL identifying the publication in the public portal

Take down policyIf you believe that this document breaches copyright please contact us providing details, and we will remove access to the work immediatelyand investigate your claim.

Download date: 10. Apr. 2022

Page 2: Charles Darwin University The extinction risk of New

1

The extinction risk of New Zealand chondrichthyans

Brittany Finucci1, Clinton A.J. Duffy2, Malcolm P. Francis3, Claudine Gibson4, Peter M. Kyne1

1 Research Institute for the Environment and Livelihoods, Charles Darwin University,

Ellengowan Dr, Darwin, NT 0909, Australia

2 Department of Conservation, Carlaw Park, 12-16 Nicholls Ln, Parnell, Auckland 1145, New

Zealand

3 National Institute of Water and Atmospheric Research (NIWA), 301 Evans Bay Pde, Greta

Point, Wellington, 6021, New Zealand

4 Auckland Zoo, Motions Rd, Auckland 1022, New Zealand

Key words: shark, ray, chimaera, conservation, IUCN Red List of Threatened Species,

deepwater, marine management

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Abstract 1

2

1. The national extinction risk of New Zealand chondrichthyans (sharks, rays, and chimaeras), 3

which accounts for ~10% of the global chondrichthyan fauna, was evaluated for the first time 4

using the IUCN Red List of Threatened Species Categories and Criteria. Across 32 families, 103 5

species were assessed. 6

2. New Zealand holds a high degree of species endemism (20%) with deepwater species 7

dominating the fauna (77%). Sharks were the most speciose group with 68 species (66%), 8

followed by 24 rays (23%), and 11 chimaeras (10%). 9

3. Most species were assessed as Least Concern (60%, 62 species) or Data Deficient (32%, 33 10

species), with four (3.8%) species listed as Near Threatened, and four (3.8%) in a threatened 11

category (Vulnerable, Endangered, Critically Endangered). Threatened species are all oceanic 12

pelagic, of which two are only visitors to New Zealand waters, and their status the result of 13

broader regional declines. 14

4. These results are in stark contrast to other recent regional assessments in Europe and the 15

Arabian Sea and adjacent areas, where up to half of species were listed in a threatened category. 16

However, given New Zealand’s extensive deepwater fishing effort and rapid collapses of 17

deepwater chondrichthyan fisheries elsewhere, it is possible that New Zealand populations of 18

many deepwater species are the remnants of previously reduced populations which are now at a 19

low, yet stable level. Ongoing species-level catch monitoring will be required to ensure these 20

species do not become threatened. 21

5. Recommendations for future research and conservation efforts include resolving taxonomic 22

uncertainties, understanding habitat use, and increasing regional collaborations to better 23

understand the effects of fishing on wider-ranging species. 24

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1. Introduction 25

26

Sharks, rays, and chimaeras (Class Chondrichthyes) are under increasing global threat. 27

Their general life history features (e.g. late maturity, slow growth, low fecundity) which result in 28

low biological productivity, as well as demand for shark products in domestic and international 29

markets, increase their susceptibility to overfishing (Daley, Stevens, & Graham, 2002; Dulvy et 30

al., 2014). Population declines have been recorded across a number of regions and species (e.g. 31

Graham, Andrew, & Hodgson, 2001; Jabado et al., 2017), with some species disappearing from 32

areas altogether (e.g. Luiz & Edwards, 2011; Dulvy et al., 2016). 33

One hundred and twelve chondrichthyan species from 32 families have been reported 34

from New Zealand’s Exclusive Economic Zone (EEZ) (Ford et al., 2018). The diversity of 35

cartilaginous fishes (~10% of the global chondrichthyan fauna) includes coastal, pelagic, and 36

deepwater species, some of which are wide-ranging while others are known only from limited 37

distributions within New Zealand. With improved species identification, taxonomic resolution, 38

and further exploration of the marine environment, this diversity continues to increase. New 39

species (e.g. Kemper, Ebert, Naylor, & Didier, 2014), and new species records (e.g. Duffy, 40

Forrester, Gibson, & Hathaway, 2017) are documented regularly. 41

Commercial fishing is the primary threat to New Zealand chondrichthyans (Ford et al., 42

2018). While chondrichthyans do not make up large proportions of catches from commercial 43

fisheries, New Zealand does have a long history of shark, ray, and chimaera fisheries (see 44

Francis, 1998). Commercial fisheries for species such as school shark (Galeorhinus galeus (L., 45

1758)) began as early as the 1900s, but landings are thought to have remained low until demand 46

for flesh and liver oil rose in the 1940s and 1950s (Francis, 1998). Today. chondrichthyans are 47

often not targeted in New Zealand, but bycatch is regularly utilized both locally and 48

internationally (Clarke, Francis, & Griggs, 2013; Francis, 1998). Approximately 80 species are 49

reported in the catches of commercial vessels, with spiny dogfish (Squalus acanthias L., 1758) 50

being the most recorded species (~24 000 t recorded between 2008-13, Francis, 2015). In 51

addition, small quantities of chondrichthyans such as rig (Mustelus lenticulatus Phillipps, 1932) 52

and elephantfish (Callorhinchus milii Bory de Saint-Vincent, 1823) are taken by recreational 53

fishers, while larger species like the shortfin mako (Isurus oxyrinchus Rafinesque, 1810) have 54

been popular with big game fishers (Francis, 1998, MPI, 2013). These species, and others, also 55

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have customary significance, as chondrichthyans were an important source of food, oil, 56

jewellery, and tools for Maori communities (Francis, 1998). 57

New Zealand chondrichthyans are managed under one of the following categories to 58

ensure sustainable utilization or protection: 11 species are managed under the Quota 59

Management System (QMS), seven species are fully protected, and two species are prohibited as 60

targets under Schedule 4C of the Fisheries Act 1996 (Table 1). All other species (i.e. most New 61

Zealand chondrichthyans) are open access. These are predominately deepwater species with 62

negligible economic value. There is no species-specific management of these species and little 63

mitigation in place to protect them or to reduce catches (MPI, 2013). 64

In addition to this legislation, New Zealand has implemented several strategies to guide 65

the management and conservation of its chondrichthyan species. These have included the 66

development of its National Plan of Action for sharks (NPOA), first released in 2008, reviewed 67

in 2013, and expected to be reviewed again in 2018 (MPI, 2013). Outlined as an objective for the 68

NPOA, a qualitative (Level 1) risk assessment with a modified Scale Intensity Consequence 69

Analysis (SICA) was conducted in 2014 for all New Zealand chondrichthyans to assess risk from 70

commercial fishing, with the intention to inform management and assist prioritizing action (Ford 71

et al., 2015). This assessment was reviewed and updated for 50 taxa in 2017 (Ford et al., 2018). 72

New Zealand is a member of Regional Fisheries Bodies (RFBs), including the Western and 73

Central Pacific Fisheries Commission (WCPFC), and internationally, is a Party to the 74

Convention on Migratory Species (CMS), signatory of the Memorandum of Understanding on 75

the Conservation of Migratory Sharks (Sharks MOU, also an NPOA objective), and a Party to 76

the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES). 77

More recently, the conservation status of all known New Zealand chondrichthyan taxa was 78

reassessed using the New Zealand Threat Classification System (NZTCS) (Duffy et al., 2018), a 79

system tailored for New Zealand (Townsend et al., 2008), but which differs considerably from 80

the IUCN Red List of Threatened Species, the world’s most utilized extinction risk framework 81

(IUCN, 2012a). 82

The IUCN Red List is the most comprehensive inventory of the global status of animal, 83

plant and fungi species, employing a single standardized set of categories and criteria to evaluate 84

the extinction risk of tens of thousands of species worldwide (Collen et al., 2016; IUCN, 2012a; 85

Mace et al., 2008). The IUCN Species Survival Commission’s Shark Specialist Group (SSG) is 86

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currently undertaking a global reassessment of all chondrichthyan fishes worldwide (~1 250 87

species), which will provide the first baseline of changes in the global and regional status of 88

chondrichthyans since the original assessments conducted >10 years ago (Dulvy et al., 2014). 89

This paper aims to assess the extinction risk of New Zealand chondrichthyans at the 90

national level. Previous assessments of New Zealand species have been included within broader 91

regional or global reviews (e.g. Cavanagh, Kyne, Fowler, Musick, and Bennett, 2003). By 92

applying the IUCN Red List of Threatened Species Categories and Criteria, the following work 93

presents a comparative view of the extinction risk of New Zealand chondrichthyans relative to 94

other regions where similar regional-level assessments have been undertaken. The completion of 95

a comprehensive national assessment will complement management tools and assist in guiding 96

future conservation and research efforts. 97

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2. Methods 98

99

A preliminary species list of 112 New Zealand chondrichthyans was compiled using the 100

most recently available knowledge as presented in Fishes of New Zealand, a comprehensive 101

review of all New Zealand fishes published by the Museum of New Zealand Te Papa Tongarewa 102

(Roberts, Stewart, & Struthers, 2015). This species list was refined by removing vagrants (a 103

taxon only occasionally found within the boundaries of a region) and species with unresolved 104

taxonomy, including formally undescribed species, to produce a final list of 103 species. Species 105

were assessed as breeding (reproduces within the region, which may involve the entire 106

reproductive cycle or any essential part of it), or visiting (does not reproduce within a region but 107

regularly occurs within its boundaries) populations (IUCN, 2012b). All available information, 108

including published reports (e.g. fisheries-independent research surveys, national fisheries stock 109

assessments, indicator analyses, technical reports), government documents (e.g. National Plan of 110

Action-Sharks, risk assessments), relevant scientific journal publications, and unpublished 111

literature was compiled for each species. 112

A two-day workshop was carried out by five experts and members of the IUCN Species 113

Survival Commission’s Shark Specialist Group (SSG). Each chondrichthyan species was 114

assessed against the IUCN Red List Categories and Criteria (Version 3.1) (IUCN, 2012a; IUCN 115

Standards and Petitions Subcommittee, 2017), together with the Guidelines for Application of 116

IUCN Red List Criteria at Regional and National Levels (Version 4.0) (IUCN, 2012b). The 117

regional guidelines can be applied at “any sub-global geographically-defined area, such as a 118

continent, country, state, or province” (IUCN, 2012b). In this study, guidelines were applied at 119

the national level, and assessments were undertaken using population sizes and trends, threats, 120

and extinction risk only within the New Zealand EEZ. For New Zealand endemic species, the 121

national assessment by default became the global assessment (since the species is found nowhere 122

else), and as such these Red List assessments were submitted to IUCN for publication on the Red 123

List of Threatened Species (http://www.iucnredlist.org/). For those species assessed as visiting, 124

the current status of the source population was evaluated, and the species’ New Zealand status 125

was assessed to be consistent with that of regional populations. 126

Each species was assessed against each of five quantitative criteria, referred to as criteria 127

A-E: Criterion A, population size reduction; B, geographic range; C, small population size and 128

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decline; D, very small or restricted population; and E, quantitative analysis (for example, a 129

population viability analysis indicating a probability of extinction; these are not available for any 130

New Zealand chondrichthyan). Species were assigned to one of the following IUCN Red List 131

categories: Extinct (EX), Extinct in the Wild (EW), Critically Endangered (CR), Endangered 132

(EN), Vulnerable (VU), (collectively, CR, EN, and VU are the ‘threatened’ categories), Near 133

Threatened (NT), Least Concern (LC), or Data Deficient (DD) (for definitions, see IUCN, 134

2012a). The DD category is applied to taxa where there is inadequate information available to 135

make an assessment of extinction risk (IUCN, 2012a). If a species qualified for a change in status 136

from a previously published assessment (a ‘down-listing’ or ‘up-listing’ in status), changes were 137

classified as genuine (a change in its extinction risk), or non-genuine (due to new information, or 138

an error in the previous assessment) (IUCN Standards and Petitions Subcommittee, 2017). 139

Following the regional IUCN guidelines, an evidentiary attitude was considered, and a species 140

was listed as threatened only when evidence to support a threatened classification was presented. 141

See IUCN (2012a) and IUCN Standards and Petitions Subcommittee (2017) for explanations and 142

guidelines of use. 143

To evaluate the diversity and degree of threat across chondrichthyan groups, species were 144

divided into subgroups based on their geographic occurrence [New Zealand endemic, Australasia 145

(New Zealand, Australia, and neighbouring Pacific nations), wider Pacific Ocean, global (wide-146

ranging across multiple ocean basins)], and major habitat and depth associations [coastal: 147

primary distribution from the shore to 12 nautical miles, nm (New Zealand’s Territorial Sea); 148

pelagic: primary distribution beyond 12 nm and at depths <200 m; deepwater: primary 149

distribution at depths >200 m). Geographic occurrence and habitat association data were largely 150

compiled from Last and Stevens (2009), Roberts et al. (2015), and Last et al. (2016). The New 151

Zealand regional assessment was compared with regional chondrichthyan extinction risk 152

assessments, which have been conducted recently for Europe (Nieto et al., 2015) and the Arabian 153

Sea and adjacent areas (hereinafter referred to as Arabian Sea, Jabado et al., 2017).154

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3. Results 155

156

Species Diversity and Endemism 157

A total of 103 species across 32 families were considered for assessment (Table 2). 158

Sharks were the most speciose group with 68 species from 22 families (66%), followed by 24 159

rays from seven families (23%), and 11 chimaeras from three families (10%). Twenty percent of 160

species were endemic to New Zealand waters, with rays and chimaeras having high levels of 161

endemism at 38% and 36%, respectively, while shark endemism was 12%. Half of the species 162

assessed from New Zealand are globally distributed, 17% are restricted to Australasia, and 12% 163

to the greater Pacific region. Most chimaeras (>70%) were restricted to New Zealand or 164

Australasian waters, while most sharks (64%) were distributed globally. The majority of species 165

(77%) were classified as deepwater, with chimaeras having the highest proportion of deepwater 166

species (91%). Across all chondrichthyans, 17% and 8% of species were classified as pelagic and 167

coastal, respectively (Table 2). 168

169

Extinction risk 170

Most species were assessed as either Least Concern (60%, 62 species) or Data Deficient 171

(32%, 33 species) with four (3.8%) listed as Near Threatened, two (1.9%) as Vulnerable, and one 172

each as Endangered (<1%) and Critically Endangered (<1%) (Table 3). Across groups, 173

chimaeras were primarily LC (91%, 10 species) with one DD species, leopard chimaera 174

(Chimaera panthera Didier, 1998). Rays were equally assessed as LC or DD, and sharks were 175

mostly LC (56%), followed by DD (32%), NT (6%), VU (3%), EN (1.5%), and CR (1.5%) 176

(Figure 2). Twelve families were found to have high levels of data deficiency (>30% of species 177

within the family assessed as DD). Arhynchobatidae and Somniosidae were the most speciose 178

families with high levels of data deficiency, with 77% and 50%, respectively (Table 4). 179

180

Threatened breeding species 181

Two species (1.9%) were listed as VU: white shark (Carcharodon carcharias L., 1758,) 182

and basking shark (Cetorhinus maximus (Gunnerus, 1765)). Both species are now protected in 183

New Zealand, but are still caught incidentally in trawl and set net fisheries (Francis & Lyon, 184

2012a, Francis, 2017a, Francis, 2017b). The total population of white sharks on the east coast of 185

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Australia and New Zealand was recently estimated to be 5460 individuals, including 750 mature 186

individuals (Bruce et al., 2018), suggesting the population within New Zealand waters alone met 187

Criterion D (‘very small or restricted population’, <1000 mature individuals). Given their 188

naturally low population size, combined with a low estimate of maximum intrinsic rate of 189

population increase (rmax) (Pardo, Kindsvater, Reynolds, & Dulvy, 2016), and documented 190

interactions with fishers, some continuing population decline is projected (10% over three 191

generations; generation length = 39 years based on age data from Natanson and Skomal, 2015), 192

the white shark also met Criterion C (‘small population size and decline’). 193

Basking shark met the criteria for VU under Criterion A (‘population reduction measured 194

over the longer of 10 years or three generations’) and Criterion C. Observed raw catch per unit 195

effort (CPUE) by trawlers showed that peak abundance of basking shark occurred in 1988-91, 196

corresponding with Japanese vessels catching relatively large numbers (>50) in some years 197

(Francis & Sutton, 2012). It is unknown if basking shark was targeted for liver oil and fins, or if 198

there was a high abundance of sharks during this period, however, catch has been near or at zero 199

since the mid-2000s, which may reflect a change in fishing gear, regional availability of sharks, 200

or a true decline in abundance (Francis, 2017b). A total of 922 individuals were estimated to 201

have been taken as commercial bycatch from 1994-95 to 2007-08, although this estimate does 202

not include captures in unobserved set net and inshore trawl fisheries (Francis & Smith, 2010). 203

This level of catch is comparable to that which took place from 1945 to 1970 off the coast of 204

British Columbia, Canada, where an estimated minimum population size of 750 was reduced to a 205

near local extinction (COSEWIC, 2007). Population trend therefore met Criterion A, that is, a 206

suspected population decline of >30% over the past three generations [generation length 207

estimated as 34 years, Pauly (1978) and Compagno (1984)], due to ‘actual levels of 208

exploitation’. Furthermore, population size within New Zealand is likely to be <10 000 mature 209

individuals, with a projected continuing decline over three generations (Criterion C) based upon 210

estimates of global population size and trend (e.g. Westgate, Koopman, Siders, Wong, & 211

Ronconi, 2014). 212

213

Near Threatened breeding species 214

Two species (1.9%) were listed as NT. Plunket shark (Scymnodon plunketi (Waite, 215

1910)) and prickly dogfish (Oxynotus bruniensis (Ogilby, 1893)) were close to meeting the 216

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criteria for the threatened categories, and were thus listed as NT. Both species have life histories 217

suggestive of low productivity (Finucci, Bustamante, Jones, & Dunn, 2016; Francis, Jones, Ó 218

Maolagáin, & Lyon, 2018), as well as high distribution overlap with fishing across much of their 219

range (45-60% and > 60% overlap, respectively) (Ford et al., 2015, 2018). While research trawl 220

survey relative biomass showed no trends in the Fisheries Management Areas (FMAs) where 221

plunket shark and prickly dogfish have been caught, monitoring of the species is poor 222

(coefficient of variation, CV, of biomass estimates >40% or greater), and, at least in the case of 223

plunket shark, reasons for a lack of trends is unknown (Francis, Roberts, & MacGibbon, 2016). 224

With estimated generation lengths of 20 (prickly dogfish) and 34 (plunket shark) years, it is 225

suspected that a population decline has occurred over three generations for both species given 226

fisheries overlap, but not at a level (30%) that would qualify for a threatened category. 227

228

Change in status 229

One species, smooth skate (Dipturus innominatus (Garrick & Paul, 1974)), was down-230

listed from NT in 2003 to LC in 2017. Implemented management efforts and indications that 231

declines in population have ceased since its previous assessment suggested smooth skate no 232

longer met NT due to a genuine change in status and was thus assessed as LC. 233

234

Visiting species 235

Five species were considered visitors to New Zealand: oceanic whitetip (Carcharhinus 236

longimanus (Poey, 1861)), dusky shark (Carcharhinus obscurus (Lesueur, 1818)), tiger shark 237

(Galeocerdo cuvier (Péron & Lesueur, 1822)), giant devilray (Mobula mobular (Bonnaterre, 238

1788)), and whale shark (Rhincodon typus Smith, 1828). While each species is infrequently 239

observed in New Zealand waters (e.g. 19 records of oceanic whitetip between 1996 and 2011, 240

Francis & Lyon, 2014), the source populations for these species from the wider Indo-Pacific 241

have undergone declines, which for some species have been considerable (>90%, Young et al., 242

2016). These declines are expected to affect the number of individuals that visit New Zealand 243

waters over time. Thus, to be consistent with the current status of regional populations, these 244

visiting species met the following criteria: Oceanic whitetip listed as CR, with a population 245

reduction of >80% over three generations based on data presented in Young et al. (2016). Dusky 246

shark is globally VU, but the source population for New Zealand is Australia, which is regionally 247

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NT (close to meeting VU A1bd) on account of additional fisheries management measures 248

implemented in 2006 (Musick, Grubbs, Baum, & Cortés, 2009); dusky shark was thus listed as 249

NT. Tiger shark was listed as NT (close to meeting criterion A2 for VU) (Simpfendorfer, 2009), 250

and whale shark as EN (past and future population reduction of >50% over three generations) 251

based on the assessment for the source population in the Indo-Pacific (Pierce & Norman, 2016). 252

Some declines in catch of Mobula spp. have been reported in fisheries managed under the 253

Western and Central Pacific Fisheries Commission (Tremblay-Boyer & Brouwer, 2016), but it is 254

not possible to determine species-specific trends in abundance since species are listed under an 255

aggregate code. Therefore, giant devilray was listed as DD at this time as the effects of fishing 256

on the species within New Zealand and in regional waters are unknown. In addition, there is 257

some evidence to suggest that the species may in fact breed within New Zealand (Duffy and 258

Tindale, 2018). 259

260

Regional comparison 261

When compared regionally, endemism in New Zealand (20%) was higher than in Europe 262

(15.2%; Nieto et al., 2015) and comparable to that of the Arabian Sea region (19.6%; Jabado et 263

al., 2017). New Zealand had a much lower proportion of threatened species (4%) than Europe 264

(32%; Nieto et al., 2015) and the Arabian Sea region (51%; Jabado et al., 2017). New Zealand 265

had the highest proportion of both DD and LC species, but unlike the other regions, had no 266

breeding species listed as EN or CR (Table 5). Forty-two species assessed in New Zealand were 267

also assessed in Europe or the Arabian Sea, 17 of which were assessed across all three regions. 268

Only one species, pelagic stingray (Pteroplatytrygon violacea (Bonaparte, 1832)) was assessed 269

as LC across all three regions. No species had a higher threat assessment in New Zealand than 270

Europe or the Arabian Sea. For some species, the extinction risk was much greater in the other 271

regions [e.g. leafscale gulper shark (Centrophorus squamosus (Bonnaterre, 1788)), LC in New 272

Zealand; EN in other regions] (Fig. 3, Appendix I). 273

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4. Discussion 274

275

New Zealand waters contain ~10% of the global chondrichthyan diversity, with a low 276

overall risk of extinction. Many species are endemic to the region (20%), and New Zealand also 277

hosts a substantial proportion of the recognized chimaeroid diversity (~20%). The low extinction 278

risk of the region (4%) is a stark contrast to other recent regional assessments, where a third and 279

over half of species in Europe and the Arabian Sea were listed in a threatened category, 280

respectively (Jabado et al., 2017; Nieto et al., 2015). Where chondrichthyans were found across 281

regions, species were listed at much higher risk of extinction outside New Zealand, particularly 282

those that are deepwater. These contrasting scenarios between New Zealand and two other 283

regions may result from New Zealand having some of the better studied and managed 284

chondrichthyan fisheries in the world, and continued effective regional management may be 285

critical for globally threatened and near-threatened species. Alternatively, historical declines in 286

these species in New Zealand may have gone undocumented, and if so, population recovery 287

would be expected to be slow, as many species are presumed to have low productivity. 288

289

New Zealand chondrichthyan management under the QMS 290

On a global scale, several New Zealand Quota Management System (QMS) 291

chondrichthyans, including elephantfish, pale ghost shark (Hydrolagus bemisi Didier, 2002), and 292

school shark, have been recognized as some of the more sustainable and well managed shark 293

fisheries (Simpfendorfer & Dulvy, 2017). Sustainable management actions have been reflected 294

in these species’ extinction risk. School shark, for example, was assessed as LC in New Zealand, 295

where relative CPUE biomass indices have been increasing or remained stable (between 1990/91 296

to 2013/14 fishing seasons) (MPI, 2017). Globally, the species is listed as VU, with considerable 297

suspected population decline in part of its range due to intensive fishing and habitat degradation 298

(Walker et al., 2006). In neighbouring Australia, school shark is listed as Conservation 299

Dependent, and targeted fishing in the Southern and Eastern Scalefish and Shark Fishery 300

(SESSF) is prohibited (AFMA, 2013). Available data suggest other species, such as blue shark 301

(Prionace glauca (L., 1758)) and shortfin mako, are increasing in abundance in New Zealand 302

(Francis, Clarke, Griggs, & Hoyle, 2014), and were listed nationally as LC. Elsewhere, these 303

species have been assessed to have a much higher extinction risk (e.g. both are listed as CR in 304

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the Mediterranean, Walls & Soldo, 2016). In the North Atlantic, the mako is considered 305

overfished (Sims, Mucientes, & Queiroz, 2018). 306

The implementation of species-specific management in New Zealand has been shown to 307

improve species status. For example, smooth skate, originally assessed as NT in 2003, was 308

down-listed to LC as a result of active management. While not targeted, this species has been, 309

and continues to be commonly caught as bycatch by benthic trawlers and longliners throughout 310

New Zealand waters (MPI, 2017). In 2002, it was highlighted that a combination of fishing 311

activities, small latitudinal range and limited depth refuge, as well as life history traits (large 312

body size; longevity >24 years; late age at maturity, 13 years for females) may threaten this 313

species (Dulvy & Reynolds, 2002). Smooth skate catches were regularly lumped together with 314

the similar looking rough skate (Zearaja nasuta (Müller & Henle, 1941)), making it impossible 315

to accurately quantify catches to the species level. In addition, landings for smooth skate had 316

exceeded quota every year since the introduction of quotas in the 1991-1992 fishing season off 317

the east coast of South Island, where most catches were recorded (Francis, 2003). Without 318

management measures to adequately regulate fishing mortality at a sustainable level, declines 319

were expected to continue, and the smooth skate was listed as NT (Francis, 2003). 320

In the same year that it was assessed, the smooth skate was introduced into the QMS, 321

along with the rough skate. The 2003 assessment of the species suggested a review of the 322

species’ status after its QMS introduction was operational and CPUE data indicated a stable 323

population. Since then, more sustainable catch limits were set, identification of skate catch 324

improved and lumped recording of the two species reduced, and live release of smooth skate 325

catch has been encouraged with its inclusion under Schedule 6 of the Fisheries Act 1996 (MPI, 326

2017, Table 1). Relative biomass estimates of smooth skate have increased with each fisheries-327

independent survey, with the 2015 estimate the highest in the time series (MPI, 2017). New 328

Zealand’s inclusion of chondrichthyans within the QMS with annually reviewed catch limits can 329

be viewed as a successful example of sustainable shark management. Catches of all QMS 330

chondrichthyans are currently at levels not considered to be overfished (MPI, 2017). These 331

efforts, however, cover only a small fraction of New Zealand’s chondrichthyan species. 332

333

The need for improved deepwater species monitoring 334

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Most New Zealand chondrichthyans (~80%) have no species-specific management or 335

monitoring. These are predominately deepwater species and are reported as bycatch in 336

commercial fisheries (Francis, 2015). At this time, these species were largely assessed as LC as 337

there was no available information to suggest any population decline or increased extinction risk. 338

Notwithstanding their sensitive life histories (Simpfendorfer & Kyne, 2009), deepwater species 339

are generally less threatened than their shallow water relatives because their distributions extend 340

beyond the depth range of most fishing effort (Dulvy et al., 2014). Without sufficient 341

management and monitoring, however, deepwater chondrichthyan populations can rapidly 342

decline from exploitation. Targeted fishing and incidental bycatch have depleted deepwater 343

chondrichthyan populations in places such as Australia (Graham et al., 2001) and the North 344

Atlantic (ICES, 2009), resulting in management arrangements such as the implementation of 345

zero catch limits (Villasante et al., 2012), and scientific recommendations to cease fishing 346

beyond 600 m (Clarke, Milligan, Bailey, & Neat, 2015). 347

In the absence of species-specific information, there is uncertainty about whether, and to 348

what degree, changes in the abundance of New Zealand chondrichthyan species have occurred 349

over time. Catch histories, which may indicate change in abundance over time when scaled with 350

fishing effort, are difficult, if not impossible, to construct for deepwater chondrichthyan bycatch 351

species in New Zealand. When recorded, catches were often aggregated under a generic code, 352

such as “deepwater dogfish” or “other sharks and dogfish” (Parker & Francis, 2012). While 353

species identification has improved over time (Francis & Lyon, 2012b), fisheries-independent 354

research trawl surveys, which assess the status of commercially valuable species on Chatham 355

Rise and the Campbell Plateau (Bagley, Ballara, O’Driscoll, Fu, & Lyon, 2013; O’Driscoll, 356

MacGibbon, Fu, Lyon, & Stevens, 2011), are the only sources of data for most deepwater 357

chondrichthyans in New Zealand. Abundance indices from these data suggest there have been 358

few trends (no change) in relative biomass for some species since the early 1990s, however, 359

monitoring for most species is poor (Ford et al., 2018; Francis et al., 2016). 360

It is possible that the abundance indices observed today in New Zealand reflect 361

previously depleted populations which are now at a low, yet stable level. Deepwater fisheries in 362

New Zealand emerged in the late 1970s, and over time, these fisheries have dominated global 363

catches of some commercial species found on the continental slope and seamounts, such as hoki 364

(Macruronus novaezelandiae (Hector, 1871)) and orange roughy (Hoplostethus atlanticus 365

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Collett, 1889), with current total allowable commercial catches (TACCs) of 150 000 t and 9800 366

t, respectively (Clarke, 2009, MPI, 2017). At least 77 chondrichthyans are reported as bycatch 367

from commercial fisheries (Anderson, 2017; Francis, 2015). Considerable declines of some 368

bycatch species associated with these deepwater fisheries has been observed. For example, over 369

a ten year period, biomass of plunket shark on north-east Chatham Rise was reported to have 370

declined in 1994 to 6% of that in the previous decade (Clark, Anderson, Francis, & Tracey, 371

2000). It is unknown how representative this area is of the entire range of plunket shark in New 372

Zealand, however, for many deepwater species there is high spatial overlap of fishing in New 373

Zealand waters (Black & Tilney, 2015; Ford et al., 2018). Eighty-five percent of trawled fishing 374

effort in New Zealand occurs at depths <800 m (Black & Tilney, 2015), overlapping with peak 375

depth distributions (Anderson et al., 1998). On Chatham Rise, New Zealand’s most productive 376

fishing ground, hoki and orange roughy stocks are considered sustainable, currently managed at 377

targets of 35-50% and 30-40% of unexploited biomass, respectively (MPI, 2017). These targets 378

may be considered desirable to achieve maximum commercial yield, however, declines of this 379

nature for deepwater chondrichthyan species with low productivity and no species-specific 380

management measures could meet the criteria for threatened categories. 381

With their generally low productivity, deepwater species are less likely to recover from 382

exploitation, and it may take decades to observe any signs of recovery towards unexploited 383

biomass levels, particularly with limited monitoring (Simpfendorfer & Kyne, 2009). Species 384

such as leafscale gulper shark and seal shark (Dalatias licha (Bonnaterre, 1788)), which are 385

regularly caught in New Zealand, have been assessed as EN in other regions on account of 386

documented declines and limited recovery (Nieto et al., 2015; Jabado et al., 2017). While this is 387

a cause for concern, under current fishing effort and management arrangements in New Zealand, 388

as well as a lack of new information since the completion of previous assessments to suggest 389

declines in populations, these species were listed as LC. This does, however, come with the 390

caveat that ongoing species-level catch monitoring is required to ensure they do not become 391

threatened. 392

393

Knowledge gaps and future efforts 394

395

Taxonomic resolution for improved fisheries reporting 396

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For many species, regardless of their assessment, there was a lack of species-specific 397

data. Of those species listed as DD, nearly all (91%) were deepwater and included many skate 398

species. There is a need for improved taxonomic resolution of morphologically conservative 399

species (e.g. Brochiraja spp.) as concerns were raised over the accuracy of catch records and 400

there was uncertainty in assessing the degree of threat for these species. Additional species (e.g. 401

members of the catshark genus Apristurus) could not be assessed at this time because of 402

taxonomic uncertainty and the use of generic codes in fishery reporting. Some of these species 403

were previously identified as having a high risk from fishing by Ford et al. (2015), with little 404

knowledge of life histories, habitat use, movement and connectivity, and population size or 405

structure. Despite having consistent, and sometimes, considerable catch records for some of 406

these species, it is difficult to assess how fishing pressures are truly affecting these populations. 407

408

Understanding habitat use and conserving habitats of importance 409

Identification and protection of habitats of importance should be investigated and 410

implemented to conserve the diversity of New Zealand’s chondrichthyans. Spatial use of the 411

water column or depth preference of different life history stages, which may increase 412

vulnerability to fishing mortality from multiple types of fishing gear (Speed, Field, Meekan, & 413

Bradshaw, 2010), is also not well known, and thus, cannot be taken into account for assessments. 414

Without an understanding of habitat use, it is difficult to assess if these areas provide any refuge 415

for most New Zealand chondrichthyans. 416

New Zealand has a series of Benthic Protected Areas (BPAs), seamount closures, and 417

marine reserve areas within its EEZ designed to manage and protect the marine environment 418

(Cryer, Mace, & Sullivan, 2016). BPAs, which protect seabed habitats through the prohibition of 419

benthic trawling and dredging, are unlikely to provide any major refuge for most 420

chondrichthyans given the distribution of species and spatial occurrence of the BPAs themselves 421

(see Black & Tilney, 2015). New industries, such as deepwater mining, should also be closely 422

monitored as they will most likely impact the New Zealand deepwater environment and benthic-423

associated species (Leduc, Rowden, Torres, Nodder, & Pallentin, 2015). The current known 424

range of one species, the Kermadec Spiny Dogfish (Squalus raoulensis Duffy & Last, 2007), lies 425

exclusively within the Kermadec Islands Marine Reserve, where fishing and mining is currently 426

prohibited. The species may also occur beyond the reserve on nearby unexplored habitat located 427

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within a designated BPA. Of all chondrichthyans, it is the only species with its known range 428

entirely within a marine protected area (Davidson & Dulvy, 2017). Virtually nothing is known 429

about the biology of this species, however, many Squalus spp. are known for their low biological 430

productivity (Graham et al., 2001). Although unlikely under the current political environment, 431

opening this area to exploitative activities may result in rapid depletion of the species. 432

433

Increased collaborations 434

With limited budgets for fisheries research, priority is given to monitoring and assessing 435

high value fish stocks, while species with little or no commercial value, such as many 436

chondrichthyans, receive little, if any, research attention (Mace, Sullivan, & Cryer, 2014). As 437

research surveys are generally the only means of data collection for many species, the collection 438

of detailed life history information beyond the usual standard length, mass, and sex 439

measurements (O’Driscoll et al., 2011), are needed to adequately describe species knowledge 440

gaps. Increased collaboration with the fishing industry and the government-sponsored observer 441

program is encouraged, as it can also allow for greater sampling coverage beyond the spatial and 442

temporal range of research surveys. 443

As some species migrate beyond New Zealand waters, including species with heightened 444

threat statuses regionally and globally, continued collaboration with neighbouring nations is 445

crucial, particularly for those species susceptible to capture in wider Pacific fisheries. For 446

example, tagging studies have indicated that giant devilray migrate seasonally between New 447

Zealand and northern subtropical or tropical areas (Francis & Jones, 2017). Catches of mobulids 448

are high outside New Zealand, however, species reporting is aggregated (Tremblay-Boyer & 449

Brouwer, 2016), limiting population monitoring. While there is little to suggest there have been 450

changes in catch rates of giant devilray within New Zealand waters, the impact of fishing outside 451

the New Zealand EEZ, and how that may affect the New Zealand visiting population, is 452

unknown (Francis & Jones, 2017). Species such as oceanic whitetip and whale shark, now listed 453

as CR and EN, respectively, have undergone considerable population declines in the Indo-West 454

Pacific due to intensive fishing pressure (Pierce & Norman, 2016; Young et al. 2016). These 455

species are considered low research priority for New Zealand given infrequent recordings (e.g. 456

Francis & Lyon, 2014), but any continued deterioration of their population outside New Zealand 457

waters will likely affect the number of visiting individuals observed within New Zealand. 458

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Ongoing collaborative projects across the South Pacific are underway to assess species’ mortality 459

of shortfin mako and silky shark (Carcharhinus falciformis (Müller & Henle, 1839)) in pelagic 460

longline fisheries (Western and Central Pacific Fisheries Commission unpub. data), and such 461

efforts could extend to other species in the future. 462

463

The national risk of extinction for New Zealand chondrichthyans has been assessed, and 464

overall, it is concluded that there is a low proportion of threatened species. When compared to 465

other assessed regions, the low risk of extinction suggests that New Zealand chondrichthyans are 466

generally well managed. However, there is a lack of species-specific data and species-specific 467

management for most species, which can impede assessing the degree of threat, past or present, 468

with certainty. Increased monitoring is highly recommended to improve knowledge and ensure 469

changes in species status, resulting from management measures or resolution in species 470

identification, can be accurately assessed. 471

472

Acknowledgements 473

474

The authors would like to thank the Auckland Zoo Conservation Fund, Oceania Chondrichthyan 475

Society, WWF, and the IUCN Species Survival Commission Shark Specialist Group for 476

supporting this project. Elephantfish vector created by Tony Ayling (vectorized by Milton Tan 477

with no additional modifications, https://creativecommons.org/licenses/by-sa/3.0/). Additional 478

thanks to Auckland Zoo and staff for hosting the project workshop. Thanks to Nicholas Dulvy, 479

Rima Jabado, Riley Pollom, Colin Simpfendorfer, and William White for additional reviews of 480

Red List assessments, and to Caroline Pollock for providing Red List support. Thanks to 481

Nicholas Dulvy and Colin Simpfendorfer for constructive comments on the manuscript. PMK 482

was supported by the Shark Conservation Fund, and the Marine Biodiversity Hub, a 483

collaborative partnership supported through funding from the Australian Government’s National 484

Environmental Science Program. 485

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Roberts, C.D., Stewart, A.L., & Struthers, C.D. (2015). The Fishes of New Zealand. Wellington: 669

Te Papa Press. 670

Simpfendorfer, C. (2009). Galeocerdo cuvier. The IUCN Red List of Threatened Species 2009: 671

e.T39378A10220026. http://www.iucnredlist.org [12 July 2018]. 672

Simpfendorfer, C.A., & Kyne, P.M. (2009). Limited potential to recover from overfishing raises 673

concerns for deep-sea sharks, rays and chimaeras. Environmental Conservation, 36, 97-103. 674

Simpfendorfer, C.A., & Dulvy, N.K. (2017). Bright spots of sustainable shark fishing. Current 675

Biology, 27, R97-R98. 676

Sims, D.W., Mucientes, G. & Queiroz, N. (2018). Shortfin mako sharks threatened by 677

inaction. Science, 359, 1342-1342. 678

Speed, C.W., Field, I.C., Meekan, M.G., & Bradshaw, C.J. (2010). Complexities of coastal shark 679

movements and their implications for management. Marine Ecology Progress Series, 408, 275-680

293. 681

Townsend, A.J., de Lange, P.J., Duffy, C.A.J., Miskelly, C.M., Molloy, J., & Norton, D.A. 682

(2008). New Zealand Threat Classification System manual. Wellington: Department of 683

Conservation. 684

Tremblay-Boyer, L., & Brouwer, S. (2016). Review of available information on non-key shark 685

species including mobulids and fisheries interactions. Bali, Indonesia: Twelfth Regular Session 686

of the Scientific Committee to the Western Central Pacific Fisheries Commission. 687

Villasante, S., Morato, T., Rodriguez-Gonzalez, D., Antelo, M., Österblom, H., Watling, 688

L.…Macho, G. (2012). Sustainability of deep-sea fish species under the European Union 689

Common Fisheries Policy. Ocean & Coastal Management, 70, 31-37. 690

Walls, R.H.L. & Soldo, A. (2016). Isurus oxyrinchus. The IUCN Red List of Threatened Species 691

2016: e.T39341A16527941. http://www.iucnredlist.org [3 June 2017]. 692

Walker, T.I., Cavanagh, R.D., Stevens, J.D., Carlisle, A.B., Chiaramonte, G.E., Domingo, A., … 693

Vooren, C.M. (2006). Galeorhinus galeus. The IUCN Red List of Threatened Species 2006: 694

e.T39352A10212764. http://www.iucnredlist.org [3 June 2017]. 695

Westgate, A.J., Koopman, H.N., Siders, Z.A., Wong, S.N., & Ronconi, R.A. (2014). Population 696

density and abundance of basking sharks Cetorhinus maximus in the lower Bay of Fundy, 697

Canada. Endangered Species Research, 23, 177-185. 698

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White, W.T., Corrigan, S., Yang, L., Henderson, A.C., Bazinet, A.L., Swofford, D.L., & Naylor, 699

G.J.P. (2018). Phylogeny of the manta and devilrays (Chondrichthyes: Mobulidae), with an 700

updated taxonomic arrangement for the family. Zoological Journal of the Linnean Society, 182, 701

50-75. 702

Young, C.N., Carlson, J., Hutchinson, M., Hutt, C., Kobayashi, D., McCandless, C.T. & Wraith, 703

J. (2016). Status review report: oceanic whitetip shark (Carcharhinus longimanus). Final Report 704

to the National Marine Fisheries Service, Office of Protected Resources. Silver Spring: National 705

Oceanic and Atmospheric Administration. 706

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FIGURE LEGN EDS 707

FIGURE LEGENDS 708

709

Figure 1. Map of New Zealand showing the boundary of its Exclusive Economic Zone (EEZ), 710

North Island (NI), South Island (SI) and Kermadec Ridge, major plateaus, and the 1000 m 711

isobath (grey line). 712

713

Figure 2. By species group, proportion of New Zealand chondrichthyans in each of the IUCN 714

Red List of Threatened Species categories. 715

716

Figure 3.Of the New Zealand species (n = 42) which have also been assessed in the Arabian Sea 717

(n = 20), and Europe (n = 39), the proportion of species assessed each of the IUCN Red List of 718

Threatened Species categories by region (DD, Data Deficient; LC, Least Concern; NT, Near 719

Threatened; VU, Vulnerable; EN, Endangered; CR, Critically Endangered). 720

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Table 1. Management categories for chondrichthyans in New Zealand (MPI, 2013). 721

Management Category Quota Management System

(QMS)

Fisheries Act 1996

Protected

Schedule 7A Wildlife Act

1953

Schedule 4C

Fisheries Act 1996

Open Access

Fisheries Act 1996

Management action Individual transferable quotas

(ITQs)

No utilization permitted No target fishing; may only be

taken as bycatch

No species-specific measures

Species Elephantfish

(Callorhinchus milii

Bory de St Vincent, 1823)

Oceanic whitetip

(Carcharhinus longimanus

(Poey, 1861))†

Sharpnose sevengill shark

(Heptranchias perlo

(Bonnaterre, 1788))

All other species

Smooth skate

(Dipturus innominatus

(Garrick & Paul, 1974))‡

White shark

(Carcharodon carcharias (L.,

1758))†

Smooth hammerhead

(Sphyrna zygaena (L., 1758))

School shark

(Galeorhinus galeus (L.,

1758))‡

Basking shark

(Cetorhinus maximus

(Gunnerus, 1765))†

Pale ghost shark

(Hydrolagus bemisi Didier,

2002

Giant manta ray

(Mobula birostris (Walbaum,

1792))

Dark ghost shark

(Hydrolagus novaezealandiae

(Fowler, 1911))

Spinetail devilray

(Mobula japanica (Müller &

Henle, 1841))§

Shortfin mako

(Isurus oxyrinchus Rafinesque,

1810)‡

Smalltooth sand tiger

(Odontaspis ferox (Risso,

1810))

Porbeagle Whale shark

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722

(Lamna nasus (Bonnaterre,

1788))‡

(Rhincodon typus Smith, 1828)

Rig

(Mustelus lenticulatus

Phillipps, 1932)‡

Blue shark

(Prionace glauca (L., 1758))‡

Spiny dogfish

(Squalus acanthias L., 1758)‡

Rough skate

(Zearaja nasuta (Müller &

Henle, 1841))‡

†Species also protected under the Fisheries Act 1996

‡ Species also included in Schedule 6 of the Fisheries Act 1996 where catch can be returned alive if the individual is likely to survive and the return takes place as

soon as possible (blue shark, porbeagle, mako, and spiny dogfish can be returned dead or alive)

§Mobula japanica is a junior synonym of Mobula mobular (Bonnaterre, 1788) (White et al., 2018)

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Table 2. The number of New Zealand chondrichthyans assessed against the IUCN Red List of 723

Threatened Species Categories and Criteria by major taxonomic group, and the proportion of 724

each group by major geographic occurrence and habitat association. 725

726

Geographic Occurrence Habitat Association

Group n

families

n

species

Endemic Australasia Greater

Pacific

Global Coastal Pelagic Deepwater

Sharks 22 68 0.120 0.100 0.150 0.630 0.060 0.200 0.740

Rays 7 24 0.380 0.290 0.040 0.290 0.125 0.125 0.750

Chimaeras 3 11 0.365 0.365 0.090 0.180 0.090 0.000 0.910

All

Chondrichthyans

32 103 0.200 0.180 0.120 0.500 0.080 0.160 0.760

727

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Table 3. National extinction risk of all New Zealand chondrichthyans (in alphabetical order by 728

family, genus and species) assessed against the IUCN Red List of Threatened Species Categories 729

and Criteria. IUCN Red List of Threatened Species categories: DD, Data Deficient; LC, Least 730

Concern; NT, Near Threatened; VU, Vulnerable; EN, Endangered; CR, Critically Endangered. 731

See IUCN (2012a) for explanations of Categories and Criteria. 732

733

Family Species name Common name National

Red List

Assessment

Alopiidae Alopias superciliosus Bigeye thresher LC

Alopias vulpinus Thresher shark DD

Arhynchobatidae Arhynchobatis asperrimus Longtail skate† LC

Bathyraja pacifica Pacific blonde skate† LC

Bathyraja richardsoni Richardson's skate LC

Bathyraja shuntovi Longnose deepsea skate† DD

Brochiraja albilabiata Whitemouth skate† DD

Brochiraja asperula Smooth deepsea skate† DD

Brochiraja heuresa Eureka skate DD

Brochiraja leviveneta Blue deepsea skate DD

Brochiraja microspinifera Dwarf skate† DD

Brochiraja spinifera Prickly deepsea skate DD

Brochiraja vittacauda Ribbontail skate DD

Notoraja alisae Velcro skate LC

Notoraja sapphira Sapphire skate DD

Callorhinchidae Callorhinchus milii Elephantfish LC

Carcharhinidae Carcharhinus brachyurus Bronze whaler LC

Carcharhinus galapagensis Galapagos shark LC

Carcharhinus longimanus Oceanic whitetip shark‡ CR A2bd

Carcharhinus obscurus Dusky shark‡ NT

Carcharhinus plumbeus Sandbar shark DD

Galeocerdo cuvier Tiger shark‡ NT

Prionace glauca Blue shark LC

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Centrophoridae Centrophorus harrissoni Harrisson's dogfish DD

Centrophorus squamosus Leafscale gulper shark LC

Deania calcea Shovelnose dogfish LC

Deania hystricosa Rough shovelnose dogfish DD

Deania quadrispinosum Longsnout dogfish DD

Cetorhinidae Cetorhinus maximus Basking shark VU A2d; C1

Chimaeridae Chimaera carophila Brown chimaera† LC

Chimaera lignaria Giant purple chimaera LC

Chimaera panthera Leopard chimaera† DD

Hydrolagus bemisi Pale ghost shark† LC

Hydrolagus homonycteris Black ghost shark LC

Hydrolagus novaezelandiae Dark ghost shark† LC

Hydrolagus trolli Pointynose blue ghost shark LC

Chlamydoselachidae Chlamydoselachus anguineus Frilled shark LC

Dalatiidae Dalatias licha Seal shark LC

Euprotomicrus bispinatus Pygmy shark LC

Isistius brasiliensis Cookiecutter shark LC

Dasyatidae Bathytoshia brevicaudata Shorttail stingray LC

Bathytoshia lata Longtail stingray LC

Pteroplatytrygon violacea Pelagic stingray LC

Echinorhinidae Echinorhinus brucus Bramble shark DD

Echinorhinus cookei Prickly shark DD

Etmopteridae Centroscyllium kamoharai Bareskin dogfish DD

Etmopterus granulosus Baxter's dogfish LC

Etmopterus lucifer Lucifer dogfish LC

Etmopterus molleri Moller's lanternshark LC

Etmopterus pusillus Smooth lanternshark LC

Etmopterus unicolor Shortspine lanternshark LC

Etmopterus viator Traveler lanternshark LC

Hexanchidae Heptranchias perlo Sharpnose sevengill shark DD

Hexanchus griseus Bluntnose sixgill shark DD

Notorynchus cepedianus Broadnose sevengill shark LC

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Lamnidae Carcharodon carcharias White shark VU

C1+2(i,ii); D1

Isurus oxyrinchus Shortfin mako LC

Lamna nasus Porbeagle LC

Mitsukurinidae Mitsukurina owstoni Goblin shark DD

Mobulidae Mobula birostris Giant manta ray LC

Mobula mobular Giant devilray‡ DD

Myliobatidae Myliobatis tenuicaudatus New Zealand eagle ray LC

Narkidae Typhlonarke aysoni Blind electric ray† LC

Odontaspididae Odontaspis ferox Smalltooth sand tiger LC

Oxynotidae Oxynotus bruniensis Prickly dogfish NT

Pentanchidae Apristurus albisoma Grey roundfin catshark DD

Apristurus ampliceps Roundfin catshark LC

Apristurus exsanguis New Zealand catshark† LC

Apristurus garricki Garrick's catshark† LC

Apristurus melanoasper Fleshynose catshark DD

Apristurus pinguis Bulldog catshark LC

Apristurus sinensis Freckled catshark LC

Parmaturus macmillani McMillan's catshark† DD

Pseudocarchariidae Pseudocarcharias kamoharai Crocodile shark DD

Pseudotriakidae Gollum attenuatus Slender smoothhound LC

Pseudotriakis microdon False catshark DD

Rajidae Amblyraja hyperborea Thorny skate LC

Dipturus innominatus Smooth skate† LC

Zearaja nasuta Rough skate† LC

Rhincodontidae Rhincodon typus Whale shark‡ EN

A2bd+4bd

Rhinochimaeridae Harriotta haeckeli Smallspine spookfish LC

Harriotta raleighana Longnose spookfish LC

Rhinochimaera pacifica Pacific spookfish LC

Scyliorhinidae Bythaelurus dawsoni Dawson's catshark† LC

Cephaloscyllium isabellum Carpet shark† LC

Somniosidae Centroscymnus coelolepis Portuguese dogfish LC

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Centroscymnus owstonii Owston's dogfish LC

Centroselachus crepidater Longnose velvet dogfish LC

Scymnodalatias albicauda Whitetail dogfish DD

Scymnodon plunketi Largespine velvet dogfish NT

Scymnodalatias sherwoodi Sherwood's dogfish DD

Scymnodon ringens Knifetooth dogfish DD

Somniosus antarcticus Pacific sleeper shark LC

Somniosus longus Little sleeper shark DD

Zameus squamulosus Velvet dogfish LC

Sphyrnidae Sphyrna zygaena Smooth hammerhead shark LC

Squalidae Cirrhigaleus australis Southern mandarin dogfish LC

Squalus acanthias Spiny dogfish LC

Squalus griffini Northern spiny dogfish† LC

Squalus raoulensis Kermadec spiny dogfish† LC

Torpedinidae Tetronarce nobiliana Great torpedo DD

Triakidae Galeorhinus galeus School shark LC

Mustelus lenticulatus Rig† LC

†Species endemic to New Zealand

‡Species treated as a visiting population to New Zealand

734

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Table 4. Poorly-known New Zealand chondrichthyan families, where >30% of New Zealand 735

species were assessed as Data Deficient. 736

737

Family Number of species in

NZ

Proportion

listed DD

Bramble sharks

Echinorhinidae

2 1.00

Torpedo rays

Torpedinidae

1 1.00

Goblin shark

Mitsukurinidae†

1 1.00

Crocodile shark

Pseudocarchariidae†

1 1.00

Softnose skates

Arhynchobatidae

13 0.77

Cowsharks

Hexanchidae

3 0.67

Gulper sharks

Centrophoridae

5 0.60

Sleeper sharks

Somniosidae

10 0.50

False catsharks

Pseudotriakidae

2 0.50

Thresher sharks

Alopiidae

2 0.50

Devilrays

Mobulidae

2 0.50

Deepwater catsharks

Pentanchidae

8 0.38

† Denotes a globally monospecific family

738

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Table 5. The comparative regional extinction risk of chondrichthyans occurring in New Zealand 739

(this study), Europe (Nieto et al., 2015), and the Arabian Seas region (Jabado et al., 2017). % 740

threatened is the sum of the categories VU, EN and CR. 741

742

n DD LC NT VU EN CR %threatened

New

Zealand

103 0.32 0.60 0.04 0.02 0.01 0.01 0.04

Europe 131 0.20 0.37 0.11 0.08 0.13 0.11 0.32

Arabian

Seas

153 0.19 0.12 0.18 0.20 0.22 0.09 0.51

IUCN Red List of Threatened Species categories: DD, Data Deficient; LC, Least Concern; NT, 743

Near Threatened; VU, Vulnerable; EN, Endangered; CR, Critically Endangered. 744

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745

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746

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747

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Appendix I. Comparative global (from IUCN, 2017) and regional extinction risk of widely-748

distributed chondrichthyans which have been assessed for a combination of New Zealand (NZ; 749

this study), Europe (Nieto et al., 2015), and/or the Arabian Seas region (Jabado et al., 2017). 750

Extinction risk was assessed by applying the IUCN Red List of Threatened Species Categories 751

and Criteria at the regional level. 752

753

Family Species Common Name Global

Status

NZ Europe Arabian

Seas

Alopiidae Alopias superciliosus Bigeye Thresher VU LC EN EN

Alopias vulpinus Thresher Shark VU DD EN --

Arhynchobatidae Bathyraja richardsoni Richardson's Skate LC LC LC --

Carcharhinidae Carcharhinus brachyurus Bronze Whaler NT LC DD --

Carcharhinus longimanus Oceanic Whitetip VU CR EN CR

Carcharhinus obscurus Dusky Shark VU NT DD --

Carcharhinus plumbeus Sandbar Shark VU DD EN EN

Galeocerdo cuvier Tiger Shark NT NT DD VU

Prionace glauca Blue Shark NT LC NT NT

Centrophoridae Centrophorus squamosus Leafscale Gulper

Shark

VU LC EN EN

Deania calcea Shovelnose

Dogfish

LC LC EN --

Deania hystricosa Rough Shovelnose

Dogfish

DD DD DD --

Cetorhinidae Cetorhinus maximus Basking Shark VU VU EN --

Chlamydoselachidae Chlamydoselachus

anguineus

Frilled Shark LC LC LC --

Dalatiidae Dalatias licha Seal Shark NT LC EN --

Dasyatidae Bathytoshia lata Longtail Stingray LC LC -- DD

Pteroplatytrygon violacea Pelagic Stingray LC LC LC LC

Echinorhinidae Echinorhinus brucus Bramble Shark DD DD EN VU

Etmopteridae Etmopterus pusillus Smooth

Lanternshark

LC LC DD DD

Hexanchidae Heptranchias perlo Sharpnose

Sevengill Shark

NT DD DD LC

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Hexanchus griseus Bluntnose Sixgill

Shark

NT DD LC LC

Lamnidae Carcharodon carcharias White Shark VU VU CR --

Isurus oxyrinchus Shortfin Mako VU LC DD NT

Lamna nasus Porbeagle VU LC CR --

Mitsukurinidae Mitsukurina owstoni Goblin Shark LC DD LC --

Mobulidae Mobula birostris† Giant Manta Ray VU LC -- VU

Mobula mobular‡ Giant Devilray NT DD EN EN

Odontaspididae Odontaspis ferox Smalltooth Sand

Tiger

VU LC CR DD

Pentanchidae Apristurus melanoasper Fleshynose

Catshark

LC DD LC --

Pseudotriakidae Pseudotriakis microdon False Catshark LC DD DD --

Rajidae Amblyraja hyperborea Thorny Skate LC LC LC --

Rhincodontidae Rhincodon typus Whale Shark EN EN -- EN

Rhinochimaeridae Harriotta haeckeli Smallspine

Spookfish

LC LC LC --

Harriotta raleighana Longnose

Spookfish

LC LC LC --

Somniosidae Centroscymnus coelolepis Portuguese

Dogfish

NT LC EN --

Centroselachus

crepidater

Longnose Velvet

Dogfish

LC LC LC DD

Scymnodon ringens Knifetooth Dogfish DD DD LC --

Zameus squamulosus Velvet Dogfish DD LC DD DD

Sphyrnidae Sphyrna zygaena Smooth

Hammerhead

Shark

VU LC DD EN

Squalidae Squalus acanthias Spiny Dogfish VU LC EN --

Torpedinidae Tetronarce nobiliana Great Torpedo DD DD LC --

Triakidae Galeorhinus galeus School Shark VU LC VU --

†Mobula birostris is currently listed as Manta birostris in the global assessment

‡Mobula mobular is currently listed as Mobula japanica in the global assessment

IUCN Red List of Threatened Species categories: DD, Data Deficient; LC, Least Concern; NT, 754

Near Threatened; VU, Vulnerable; EN, Endangered; CR, Critically Endangered. 755

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756